688 COMMUNICATION
Late Pleistocene age, size, and paleoenvironment of a caribou antler from Haida Gwaii, British Columbia Rolf W. Mathewes, Michael Richards, and Thomas E. Reimchen
Abstract: The basal portion of a fossil caribou antler from Graham Island is the only evidence of large terrestrial vertebrates older than the Fraser (late-Wisconsin) glaciation on Haida Gwaii. This antler has been radiocarbon dated three times by different laboratories and all ages fall within the mid-Wisconsin Olympia Interglaciation (Marine Isotope Stage 3, MIS 3). We suggest that the latest date, using ultrafiltration of bone collagen, is closest to the true age at 43 200 ± 650 years BP (48 200 – 45 200 cal BP). Previous paleoecological analysis from Graham Island reconstructed a vegetation cover during MIS 3 consisting of mixed coniferous forest with nonforested openings, similar to cool subalpine forests of today. These conditions are consistent with environments that support woodland caribou and the related extinct Dawson caribou. Morphometric comparison of antlers from woodland and Dawson caribou suggest that they are more similar than previously interpreted and raise questions about the inferred differences between the mainland and island subspecies.
Key words: Haida Gwaii, caribou, fossil antler, accelerator mass spectrometry dating, paleoecology, MIS 3. Résumé : La portion basale de bois de caribou fossiles de l’île Graham est le seul indice de la présence de grands vertébrés terrestres plus ancienne que la glaciation de Fraser (Wisconsinien tardif) dans l’archipel Haida Gwaii. Le panache a été daté au carbone radioactif à trois reprises par différents laboratoires, et tous les âges tombent dans la période de l’interglaciaire d’Olympia du Wisconsinien moyen (stade isotopique marin 3, MIS 3). Nous suggérons que l’âge le plus récent, obtenu par la méthode d’ultrafiltration sur collagène osseux, est celui qui se rapproche le plus de l’âge réel, à 43 200 ± 650 ans BP (48 200 – 45 200 étal. BP). Une analyse paléoécologique antérieure de l’île Graham a reconstitué une couverture végétale durant le MIS 3 consistant en une forêt de conifères mixte avec des ouvertures non boisées, semblable aux forêts subalpines de régions fraîches actuelles. Ces conditions concordent avec des milieux qui supportent le caribou des bois et l’espèce apparentée aujourd’hui disparue du caribou de Dawson. La comparaison morphométrique de panaches de caribous des bois et de Dawson indiquerait une plus grande similitude des deux espèces qu’interprétée auparavant et soulève des questions quant aux dif- férences inférées entre les sous-espèces continentales et insulaires. [Traduit par la Rédaction]
Mots-clés : Haida Gwaii, caribou, panache fossile, datation spectrométrie de masse par accélérateur, paléoécologie, MIS 3. For personal use only.
Introduction cies Rangifer dawsoni (Seton-Thompson 1900) in honour of Dawson. It was later reduced to subspecies status (Rangifer tarandus dawsoni) Haida Gwaii (formerly Queen Charlotte Islands) is a large archi- of mountain caribou (woodland caribou) (Rangifer tarandus caribou), pelago of two main islands and many smaller ones located about its likely ancestor. Three ecotypes of mountain caribou are cur- 80 km west of mainland British Columbia on the edge of the North rently recognized in British Columbia with different distributions American continental shelf. These islands have been called the (southern mountain, northern mountain, and boreal), but all are “Canadian Galapagos” and have been the subject of much re- classified as the same subspecies (Ministry of Environment 2000). search on the biogeography and endemism of their fauna and Mitochondrial DNA analysis (Byun et al. 2002) revealed that Daw- flora as well as their glacial history (Foster 1965; review papers in son caribou are not genetically distinct from mountain caribou on Scudder and Gessler 1989; Fedje and Mathewes 2005). the mainland of British Columbia, although the attributed phys- Wigen (2005), in her review of the vertebrate fauna of Haida ical characteristics of smaller size, reduced antlers, and pale pel- Gwaii, noted that the history of caribou deserves special consid- age have long been recognized as distinctive and different when eration, likely due to the fact that the endemic form called Daw- compared to mainland caribou. These characteristics are, how- son caribou (Rangifer tarandus dawsoni) has long attracted attention ever, based on few observations and a very small sample of antlers Can. J. Earth Sci. Downloaded from www.nrcresearchpress.com by UNIV VICTORIA on 06/15/19 and speculation regarding details of its origin, physical character- and skulls (including the holotype) and a skin of the last shot istics, and extinction (Seton-Thompson 1900; Foster 1965; Banfield specimen (Banfield 1963). 1963; Byun et al. 2002). It is not known if Dawson caribou is a postglacial neoendemic The geologist G.M. Dawson brought the existence of Haida island form (Reimchen and Byun 2005), an older relict taxon, or Gwaii caribou to the attention of Ernest Seton-Thompson in 1899, even an immigrant from an unknown glacial refuge on the north- who provided a preliminary description and named the new spe- west coast. It is now widely accepted that glaciation of Haida
Received 19 September 2018. Accepted 16 December 2018. Paper handled by Associate Editor Olav B. Lian. R.W. Mathewes. Department of Biological Sciences, Simon Fraser University, 8888 University Drive, Burnaby, BC V5A 1S6, Canada. M. Richards. Department of Archaeology, Simon Fraser University, Burnaby, BC V5A 1S6, Canada. T.E. Reimchen. Department of Biology, University of Victoria, Box 3020, Victoria, BC V8W 3N5, Canada. Corresponding author: Rolf W. Mathewes (email: [email protected]). Copyright remains with the author(s) or their institution(s). Permission for reuse (free in most cases) can be obtained from RightsLink.
Can. J. Earth Sci. 56: 688–692 (2019) dx.doi.org/10.1139/cjes-2018-0246 Published at www.nrcresearchpress.com/cjes on 9 January 2019. Mathewes et al. 689
Fig. 1. Map of Graham Island showing the location of the White mens of Dawson caribou available for study is very limited (sev- Creek antler collection site and other names mentioned in the text. eral antlers and skulls and one skin, without other skeletal [Colour online.] elements), it is possible that the range of body and antler sizes for Dawson caribou may have been larger than currently accepted. Dawson caribou is the only member of the deer family in which females supposedly do not possess antlers (Cowan and Guiget 1956), which complicates any generalizations due to the very small sample size of specimens and field observations. The donation by Mr. Art Lightburn of a fossil caribou antler fragment to the Queen Charlotte Islands Museum (now Haida Gwaii Museum at Kay Llnagaay) in 1981 was a significant event, since it was the first find in a nonarcheological setting and listed by Wigen (2005) as the only definite preglacial vertebrate from Haida Gwaii. The fossil was found in sediments at the mouth of White Creek (Fig. 1), also known by its Haida name of SGidlii Gandl. Mr. Lightburn recalls only that he found the antler in 1978 “in one of the washouts at the mouth of White Creek” based on a tele- phone conversation with Mr. Lightburn by Nathalie MacFarlane on December 3, 2018. White Creek (54.0526°N, 131.88904°W) flows north on the Argonaut Plain (Fig. 1) and empties into the waters of South Beach along Dixon Entrance, where tidal influences and stream discharges create shifting conditions of erosion and depo- sition in the sand and gravel at the stream mouth. This paper outlines the history of radiocarbon dating and paleo- environmental interpretation of this antler fragment. Its age, based on the first radiocarbon date, has been variously cited (as personal communication with R.W. Mathewes) at around 40 000 years or older (Byun et al. 2002; Wigen 2005) but the actual ages have not been previously published and interpreted. Materials and methods The fossil caribou antler from White Creek represents the basal portion (burr) where it is attached to the skull and the main beam with two broken tines (brow and bez) (Fig. 2). It was originally made available to R.W. Mathewes for study and radiocarbon dat- ing by the Queen Charlotte Islands Museum in 1994 and recently again for study and redating by the Haida Gwaii Museum at Kay
For personal use only. Gwaii was less extensive than on the adjacent mainland, with a Llnagaay at Skidegate, Graham Island, where it is housed under thin local ice cap that receded earlier here than elsewhere on the accession number NH 17001. coast (Clague et al. 1982; Warner et al. 1982; Mathewes and Clague 2017). There is increasing evidence that ice-free biotic refugia ex- The first radiocarbon sample was prepared by Erle Nelson at isted along the northwest coast during the last (Fraser) glaciation Simon Fraser University, who extracted collagen and sent the on both geological and biological grounds (Heusser 1989; Heaton sample for dating by accelerator mass spectrometry (AMS) to Law- et al. 1996; Barrie et al. 2005; Carrara et al. 2007; Shafer et al. 2010; rence Livermore National Laboratory in 1994. A second sample Mathewes et al. 2015; Mathewes and Clague 2017). Details of the of tissue drilled from the interior of the antler was analyzed for extent of local ice cover and biological responses to it are still DNA at the University of Victoria by Reimchen but no DNA was uncertain, although we know that lowered sea levels during late recovered. The remainder of this sample was later sent for AMS and postglacial periods exposed much of vegetated land that was radiocarbon dating to Beta Analytic Inc. in Florida, who also de- not drowned until postglacial sea levels rose again (Fedje and termined a finite, but younger age than the original date. The Josenhans 2000; Lacourse et al. 2003), thereby reducing the isola- most recent third sample using ultrafiltration dating of solid bone tion of Haida Gwaii from the mainland during glacial periods, collagen was collected by M. Richards at Simon Fraser University which would facilitate migration by cold-adapted animals like and sent to the A.E. Lalonde AMS Laboratory at the University of caribou. Ottawa for analysis. Can. J. Earth Sci. Downloaded from www.nrcresearchpress.com by UNIV VICTORIA on 06/15/19 Fossil remains of caribou are known on Graham Island, Haida To evaluate morphometrics, we measured width and thickness Gwaii, from several postglacial (ϳ6000–1500 14C years BP) arche- at each of two positions on the fossil antler fragment (Fig. 2). ological sites such as Bluejackets Creek near Masset and Cohoe Equivalent measurements were also obtained from antlers taken Creek near Port Clements (Fig. 1). Fossil caribou bones have also from three Dawson caribou killed in the late 1800s and early 1900s been dated to ca. 13 300 cal BP (Fedje and Smith 2010) at K-1 cave on and from five woodland caribou from mainland British Columbia. northwestern Moresby Island along with grizzly and black bear All specimens are curated at the Royal British Columbia Museum, remains and other vertebrates. This unpublished report confirms formerly known as the British Columbia Provincial Museum, Vic- that caribou were once present on both Graham and Moresby toria. The four highly correlated metrics were reduced with prin- islands. cipal components analysis before plotting. Regarding the Graham Island skeletal elements, Wigen (2005) pointed out that none of these show evidence of the dwarfing Results and discussion attributed to Dawson caribou, suggesting that it may be a recent The three AMS radiocarbon ages from antler fragment NH 7001 development. However, morphometric data are not presented in are summarized in Table 1, showing the original and calibrated support of the contention that the archeological specimens are ages. Although the three finite AMS ages are different, they all fall larger than those of Dawson caribou. Since the number of speci- within Marine Isotope Stage 3 (MIS 3), the mid-Wisconsin substage
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Fig. 2. (A) Distal and (B) proximal views of the fossil caribou antler (NH 17001) from White Creek. The two measurement points for width and thickness are indicated by the double-headed arrows on Fig. 2A. The attachment point to the skull (burr) is at the base of both images. Scale bar = 5 cm. [Colour online.]
Table 1. Radiocarbon ages determined from the White Creek fossil tively with consumption of intertidal algae, which are highly caribou antler (Graham Island) from Haida Gwaii. enriched in both isotopes (Fox et al. 2018). We are currently undertaking additional isotope studies of caribou (including sul-
For personal use only. Radiocarbon age Calibrated age Isotopic (14C years BP)a (cal years BP)b Laboratory No. signature 13C/12C phur isotope measurements and compound-specific amino acid isotope measurements) to better understand this surprising iso- 32 980±180 38 500 to 37 000 BETA-317984 None 43 209±649* 48 200 to 45 200 UOC-5713 Measured: 14.6‰ tope result. 47 600±2600* 57 600 to 43 300 CAMS-12921 None Paleoenvironment during MIS 3 Note: BETA, Beta Analytic Inc., Miami, Florida; UOC, A.E. Lalonde AMS Labo- The presence of caribou during the mid-Wisconsin is consistent ratory, University of Ottawa, Ottawa, Ontario, Canada; CAMS, Center for Accel- erator Mass Spectrometry, Lawrence Livermore National Laboratory, University with a local environment interpreted as being cooler than pres- of California, Berkeley, California. ent, which supported coniferous forest with subalpine elements *Material dated is collagen extracted from solid bone, except for BETA, which such as mountain hemlock (Tsuga mertensiana) and parkland- like is collagen extracted from the interior cavity tissue. openings with wet herbaceous meadows and peatland vegetation. aError term ±1. This interpretation is based on the only MIS 3 pollen and plant b Age range at ±2 using the OxCAL4.2 Online calibration program (Bronk macrofossil record from Haida Gwaii (Warner et al. 1984) recorded Ramsey 2009) with the IntCal 13 data set (Reimer et al. 2013) rounded to the nearest 100. at the Pilot Mill site on Graham Island (Fig. 1). A photograph and ecological summary of the Pilot Mill peat bed are published in
Can. J. Earth Sci. Downloaded from www.nrcresearchpress.com by UNIV VICTORIA on 06/15/19 Mathewes (1989). The Pilot Mill peat study concluded that annual also known as the Olympia Interglaciation or Olympia Intersta- temperatures were lowered by 2 °C compared to modern and dial on the west coast (Hebda et al. 2016). Based on the latest treelines were depressed by ϳ400 m. A review of mid-Wisconsin ultrafiltration date on bone collagen, we consider the 43 200 BP environments in other localities in western Canada and Alaska age to be closest to the true age. It also overlaps the age range of generally confirms cooler climatic conditions during MIS 3 (Clague the oldest age of 47 600 BP (Table 1). The youngest age (32 980 BP) et al. 2004; Hebda et al. 2016) with the most open and largely falls outside the ranges of both older finite ages for undetermined reasons, but it was determined on tissue collected originally for treeless conditions at the base of the Pilot Mill peat and also at the 14 DNA analysis from the inside (marrow region) of the antler, rather base of Lynn Valley peat at Vancouver at >48 CkaBP(Hebda than on collagen extracted from solid bone. et al. 2016). Relative to isotopic signatures for modern caribou from coastal Mountain caribou require old-growth forests as winter habitat Alaska and Svaalbard (Zwolicki et al. 2016; Pacyna et al. 2018), our and for arboreal and terrestrial lichens as winter food, and the signatures for the Haida Gwaii fossil caribou antler (␦13C = –14.6‰, pollen record at Pilot Mill indicates a forest composed of spruce ␦15N = 9.9‰) are enriched. Such elevated isotopic values could be (Picea) and true fir (Abies) in addition to mountain hemlock, likely associated with consumption of terrestrial vegetation growing in providing the needed forest cover for mountain caribou to thrive the vicinity of seabird colonies (Zwolicki et al. 2016) or alterna- during winters.
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Fig. 3. Principal components analysis of morphometrics of Dawson Although our study was not designed to evaluate the morpholog- and woodland caribou antler beams from Haida Gwaii and mainland ical uniqueness of Dawson caribou, the metrics that we identified on British Columbia. Dawson1HG is the type specimen (British Columbia the antlers allow comment on this issue. Seton-Thompson (1900) and Provincial Museum (BCPM) label 1483), Dawson2HG was shot in 1909 subsequently Cowan and Guiget (1956) characterized the antlers (BCPM label 1486), Dawson3HG is a museum mount (BCPM label 1484), of Dawson as “poorly developed”, which accurately describes the Woodland1BC (BCPM label 1469), Woodland2BC (BCPM label 6933), antler (Dawson3HG) on the reconstructed mount widely used in Woodland3BC (BCPM label 1498), Woodland4BC (BCPM label 1837), descriptions of this caribou. Yet our data on the two additional and Woodland5BC (BCPM label 10225). All individuals are male except antlers from this taxon as well as the fossil fragment show these to for Woodland5BC, which is female. The sex of the fossil antler is be similar or larger than three of the four male woodland caribou. unknown. Consequently, we suspect that the original characterization of Dawson caribou as a small-bodied form placed too much empha- sis on the museum mount, which was the smallest of the three specimens. Accordingly, the evidence for the description of Daw- son caribou as a “dwarf” subspecies of mountain caribou should be reexamined. Acknowledgements We would like to thank Nathalie MacFarlane, Archives man- ager, and the Haida Gwaii Museum at Kay Llnagaay for providing access to the fossil antler for analysis and Mr. Art Lightburn for updating information on his fossil find. M.P.R. would also like to thank the Natural Sciences and Engineering Research Council (NSERC) of Canada for funding the OUC radiocarbon date. An NSERC Discovery Grant (A3835) to R.W.M. and an NSERC Discov- ery Grant (A2354) to T.E.R. are also gratefully acknowledged for support of Haida Gwaii research, including this study. Leslie Kennes and Darren Grocke generously provided access to the caribou specimens at the Royal British Columbia Museum and Matthew Huntley drafted the map. The comments of Olav Lian (Associate Editor) and the external reviewers Daryl Fedje and Vaughn Barrie are appreciated and improved the manuscript.
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